Journal of Plant Research

, Volume 130, Issue 5, pp 829–844 | Cite as

Elevational plant species richness patterns and their drivers across non-endemics, endemics and growth forms in the Eastern Himalaya

  • Kumar Manish
  • Maharaj K. PanditEmail author
  • Yasmeen Telwala
  • Dinesh C. Nautiyal
  • Lian Pin Koh
  • Sudha Tiwari
Regular Paper


Despite decades of research, ecologists continue to debate how spatial patterns of species richness arise across elevational gradients on the Earth. The equivocal results of these studies could emanate from variations in study design, sampling effort and data analysis. In this study, we demonstrate that the richness patterns of 2,781 (2,197 non-endemic and 584 endemic) angiosperm species along an elevational gradient of 300–5,300 m in the Eastern Himalaya are hump-shaped, spatial scale of extent (the proportion of elevational gradient studied) dependent and growth form specific. Endemics peaked at higher elevations than non-endemics across all growth forms (trees, shrubs, climbers, and herbs). Richness patterns were influenced by the proportional representation of the largest physiognomic group (herbs). We show that with increasing spatial scale of extent, the richness patterns change from a monotonic to a hump-shaped pattern and richness maxima shift toward higher elevations across all growth forms. Our investigations revealed that the combination of ambient energy (air temperature, solar radiation, and potential evapo-transpiration) and water availability (soil water content and precipitation) were the main drivers of elevational plant species richness patterns in the Himalaya. This study highlights the importance of factoring in endemism, growth forms, and spatial scale when investigating elevational gradients of plant species distributions and advances our understanding of how macroecological patterns arise.


Elevational gradient Endemic Growth forms Himalaya Macroecology Richness patterns 



KM acknowledges the support of Department of Science and Technology INSPIRE Research Fellowship, Government of India (Grant No: DST/INSPIRE Fellowship/2012/432). The financial support to MKP provided by the Ministry of Environment, Forests and Wildlife, Government of India and NHPC India vide grant No. J.12011/11/99-IA.I. and DU-DST-PURSE Grant is gratefully acknowledged. LPK was supported by the Australian Research Council. We also thank D. Dawa and R. Mehta for assistance.

Compliance with ethical standards

Human and animal rights

No formal approval is required for this study since this article does not contain any studies with human participants or animals performed by any of the authors.

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

10265_2017_946_MOESM1_ESM.pdf (722 kb)
Supplementary material 1 (PDF 722 KB)


  1. Acharya BK, Chettri B, Vijayan L (2011a) Distribution pattern of trees along an elevation gradient of Eastern Himalaya, India. Acta Oecol 37:329–336CrossRefGoogle Scholar
  2. Acharya KP, Vetaas OR, Birks HJB (2011b) Orchid species richness along Himalayan elevational gradients. J Biogeogr 38:1821–1833CrossRefGoogle Scholar
  3. Bertuzzo E, Carrara F, Mari L, Altermatt F, Rodriguez-Iturbe I, Rinaldo A (2016) Geomorphic controls on elevational gradients of species richness. Proc Natl Acad Sci USA 113:1737–1742CrossRefPubMedPubMedCentralGoogle Scholar
  4. Bhatt JP, Manish K, Pandit MK (2012) Elevational gradients in fish diversity in the Himalaya: water discharge is the key driver of distribution patterns. PLoS One 7:e46237. doi: 10.1371/journal.pone.0046237 CrossRefPubMedPubMedCentralGoogle Scholar
  5. Bhattarai KR, Vetaas OR (2003) Variation in plant species richness of different life forms along a subtropical elevation gradient in the Himalayas, east Nepal. Global Ecol Biogeogr 12:327–340CrossRefGoogle Scholar
  6. Bhattarai KR, Vetaas OR (2006) Can Rapoport’s rule explain tree species richness along the Himalayan elevation gradient, Nepal? Divers Distrib 12:373–378CrossRefGoogle Scholar
  7. Brown JH, Lomolino MV (1998) Biogeography. Sinauer Associates, SunderlandGoogle Scholar
  8. Carpenter C (2005) The environmental control of plant species density on a Himalayan elevation gradient. J Biogeogr 32:999–1018CrossRefGoogle Scholar
  9. Chettri B, Bhupathy S, Acharya BK (2010) Distribution pattern of reptiles along an eastern Himalayan elevation gradient, India. Acta Oecol 36:16–22CrossRefGoogle Scholar
  10. Colwell RK (2013) EstimateS: statistical estimation of species richness and shared species from samples. Version 9.1. Accessed 4 May 2014
  11. Colwell RK, Hurtt GC (1994) Nonbiological gradients in species richness and a spurious Rapoport effect. Am Nat 144:570–595CrossRefGoogle Scholar
  12. Colwell RK, Lees DC (2000) The mid-domain effect: geometric constraints on the geography of species richness. Trends Ecol Evol 15:70–76CrossRefPubMedGoogle Scholar
  13. Colwell RK, Rahbek C, Gotelli NJ (2004) The mid-domain effect and species richness patterns: what have we learned so far? Am Nat 163:E1–E23CrossRefPubMedGoogle Scholar
  14. Connell JH (1978) Diversity in tropical rain forests and coral reefs. Science 199:1302–1310CrossRefPubMedGoogle Scholar
  15. Currie DJ, Kerr JT (2008) Tests of the mid-domain hypothesis: a review of the evidence. Ecol Monogr 78:3–18CrossRefGoogle Scholar
  16. da Silva FKG, de Faria Lopes S, Lopez LCS, de Melo JIM, Trovão DMDBM (2014) Patterns of species richness and conservation in the Caatinga along elevational gradients in a semiarid ecosystem. J Arid Environ 110:47–52CrossRefGoogle Scholar
  17. Field R, Hawkins BA, Cornell HV, Currie DJ, Diniz-Filho JA, Guégan JF, Kaufman DM, Kerr JT, Mittelbach GG, Oberdorff T, O’Brien EM (2009) Spatial species-richness gradients across scales: a meta-analysis. J Biogeogr 36:132–147CrossRefGoogle Scholar
  18. Francis AP, Currie DJ (2003) A globally consistent richness-climate relationship for angiosperms. Am Nat 161:523–536CrossRefPubMedGoogle Scholar
  19. Fraser LH, Pither J, Jentsch A et al (2015) Worldwide evidence of a unimodal relationship between productivity and plant species richness. Science 349:302–305CrossRefPubMedGoogle Scholar
  20. Fu C, Hua X, Li J, Chang Z, Pu Z, Chen J (2006) Elevational patterns of frog species richness and endemic richness in the Hengduan mountains, China: geometric constraints, area and climate effects. Ecography 29:919–927CrossRefGoogle Scholar
  21. Gaston KJ, Chown SL (1999) Why Rapoport’s rule does not generalise. Oikos 84:309–312CrossRefGoogle Scholar
  22. González-Oreja J, Garbisu C, Mendarte S, Ibarra A, Albizu I (2010) Assessing the performance of nonparametric estimators of species richness in meadows. Biodivers Conserv 19:1417–1436CrossRefGoogle Scholar
  23. Grace JB, Anderson TM, Seabloom EW et al (2016) Integrative modelling reveals mechanisms linking productivity and plant species richness. Nature 529:390–393CrossRefPubMedGoogle Scholar
  24. Graham JH, Duda JJ (2011) The humpbacked species richness-curve: a contingent rule for community ecology. Int. J Ecol 2011:1–15Google Scholar
  25. Grau O, Grytnes JA, Birks HJB (2007) A comparison of altitudinal species richness patterns of bryophytes with other plant groups in Nepal, Central Himalaya. J Biogeogr 34:1907–1915CrossRefGoogle Scholar
  26. Grime JP (1977) Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. Am Nat 111:1169–1194CrossRefGoogle Scholar
  27. Grime JP (2001) Plant strategies, vegetation processes, and ecosystem properties. Wiley, ChichesterGoogle Scholar
  28. Grinnell J, Storer TI (1924) Animal life in the Yosemite: an account of the mammals, birds, reptiles, and amphibians in a cross-section of the Sierra Nevada. University of California Press, BerkeleyGoogle Scholar
  29. Grytnes JA (2003) Ecological interpretations of the mid-domain effect. Ecol Lett 6:883–888CrossRefGoogle Scholar
  30. Grytnes JA, McCain CM (2007) Elevational trends in biodiversity. In: Levin S (ed) Encyclopedia of Biodiversity. Elsevier, New York, pp 1–8Google Scholar
  31. Grytnes JA, Vetaas OR (2002) Species richness and altitude: a comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient, Nepal. Am Nat 159:294–304CrossRefPubMedGoogle Scholar
  32. Grytnes JA, Heegaard E, Romdal TS (2008) Can the mass effect explain the mid-altitudinal peak in vascular plant species richness? Basic Appl Ecol 9:373–382CrossRefGoogle Scholar
  33. Harrison S, Cornell H (2008) Toward a better understanding of the regional causes of local community richness. Ecol Lett 11:969–979CrossRefPubMedGoogle Scholar
  34. Hooker JD (1875–1897) The flora of British India, vol 1–7. L. Reeve and Co., LondonGoogle Scholar
  35. Hu J, Xie F, Li C, Jiang J (2011) Elevational patterns of species richness, range and body size for spiny frogs. PLoS One 6:e19817. doi: 10.1371/journal.pone.0019817 CrossRefPubMedPubMedCentralGoogle Scholar
  36. Huston MA (2014) Disturbance, productivity, and species diversity: empiricism vs. logic in ecological theory. Ecology 95:2382–2396CrossRefGoogle Scholar
  37. Kallimanis AS, Bergmeier E, Panitsa M, Georghiou K, Delipetrou P, Dimopoulos P (2010) Biogeographical determinants for total and endemic species richness in a continental archipelago. Biodivers Conserv 19:1225–1235CrossRefGoogle Scholar
  38. Karger DN, Kluge J, Krömer T, Hemp A, Lehnert M, Kessler M (2011) The effect of area on local and regional elevational patterns of species richness. J Biogeogr 38:1177–1185CrossRefGoogle Scholar
  39. Kerr JT (2001) Butterfly species richness patterns in Canada: energy, heterogeneity, and the potential consequences of climate change. Conserv Ecol 5:10CrossRefGoogle Scholar
  40. Kessler M (2002) The elevational gradient of Andean plant endemism: varying influences of taxon-specific traits and topography at different taxonomic levels. J Biogeogr 29:1159–1165CrossRefGoogle Scholar
  41. Kharkwal G, Mehrotra P, Rawat YS, Pangtey YP (2005) Phytodiversity and growth form in relation to altitudinal gradient in the Central Himalayan (Kumaun) region of India. Curr Sci 89:873–878Google Scholar
  42. Körner C (1999) Alpine plant life: functional plant ecology of high mountain ecosystems. Springer Science & Business Media, BerlinCrossRefGoogle Scholar
  43. Körner C (2007) The use of ‘altitude’ in ecological research. Trends Ecol Evol 22:569–574CrossRefPubMedGoogle Scholar
  44. Lamoreux JF, Morrison JC, Ricketts TH, Olson DM, Dinerstein E, McKnight MW, Shugart HH (2006) Global tests of biodiversity concordance and the importance of endemism. Nature 440:212–214CrossRefPubMedGoogle Scholar
  45. Lomolino MV (2001) Elevation gradients of species-density: historical and prospective views. Global Ecol Biogeogr 10:3–13CrossRefGoogle Scholar
  46. Manish K, Telwala Y, Nautiyal DC, Pandit MK (2016) Modelling the impacts of future climate change on plant communities in the Himalaya: a case study from Eastern Himalaya, India. Model Earth Syst Environ 2:92. doi: 10.1007/s40808-016-0163-1 CrossRefGoogle Scholar
  47. McCain CM (2003) North American desert rodents: a test of the mid-domain effect in species richness. J Mammal 84:967–980CrossRefGoogle Scholar
  48. McCain CM (2004) The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica. J Biogeogr 31:19–31CrossRefGoogle Scholar
  49. McCain CM (2007a) Area and mammalian elevational diversity. Ecology 88:76–86CrossRefPubMedGoogle Scholar
  50. McCain CM (2007b) Could temperature and water availability drive elevational species richness patterns? A global case study for bats. Global Ecol Biogeogr 16:1–13CrossRefGoogle Scholar
  51. McCain CM, Knight BK (2013) Elevational Rapoport’s rule is not pervasive on mountains. Global Ecol Biogeogr 22:750–759CrossRefGoogle Scholar
  52. Nogues-Bravo D, Araujo MB, Romdal T, Rahbek C (2008) Scale effects and human impact on the elevational species richness gradients. Nature 453:216–219CrossRefPubMedGoogle Scholar
  53. Oommen MA, Shanker K (2005) Elevational species richness patterns emerge from multiple local mechanisms in Himalayan woody plants. Ecology 86:3039–3047CrossRefGoogle Scholar
  54. Orme CDL, Davies RG, Burgess M, Eigenbrod F, Pickup N, Olson VA, Webster AJ, Ding TS, Rasmussen PC, Ridgely RS, Stattersfield AJ (2005) Global hotspots of species richness are not congruent with endemism or threat. Nature 436:1016–1019CrossRefPubMedGoogle Scholar
  55. Pan X, Ding Z, Hu Y, Liang J, Wu Y, Si X, Guo M, Hu H, Jin K (2016) Elevational pattern of bird species richness and its causes along a central Himalaya gradient, China. PeerJ 4:e2636. doi: 10.7717/peerj.2636 CrossRefPubMedPubMedCentralGoogle Scholar
  56. Pandit MK (2017) Life in the Himalaya: an ecosystem at risk. Harvard University Press, CambridgeGoogle Scholar
  57. Pandit MK, Sodhi N, Koh L, Bhaskar A, Brook B (2007) Unreported yet massive deforestation driving loss of endemic biodiversity in Indian Himalaya. Biodivers Conserv 16:153–163CrossRefGoogle Scholar
  58. Pandit MK, Manish K, Koh LP (2014) Dancing on the roof of the world: ecological transformation of the Himalayan landscape. Bioscience 64:980–992CrossRefGoogle Scholar
  59. R Development Core Team (2011) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0. Accessed 22 Aug 2014
  60. Raes N, Roos MC, Slik JW, Van Loon EE, Steege HT (2009) Botanical richness and endemicity patterns of Borneo derived from species distribution models. Ecography 32:180–192CrossRefGoogle Scholar
  61. Rahbek C (2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecol Lett 8:224–239CrossRefGoogle Scholar
  62. Singh JS, Singh SP (1987) Forest vegetation of the Himalaya. Bot Rev 53:80–192CrossRefGoogle Scholar
  63. Stein A, Gerstner K, Kreft H (2014) Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales. Ecol Lett 17:866–880CrossRefPubMedGoogle Scholar
  64. Stevens GC (1992) The elevational gradient in altitudinal range: an extension of Rapoport’s latitudinal rule to altitude. Am Nat 140:893–911CrossRefPubMedGoogle Scholar
  65. Stohlgren TJ, Falkner MB, Schell LD (1995) A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113–121CrossRefGoogle Scholar
  66. Telwala Y, Brook BW, Manish K, Pandit MK (2013) Climate-induced elevational range shifts and increase in plant species richness in a Himalayan biodiversity epicentre. PLoS One 8:e57103. doi: 10.1371/journal.pone.0057103 CrossRefPubMedPubMedCentralGoogle Scholar
  67. Tredennick AT, Adler PB, Grace JB et al (2016) Comment on “Worldwide evidence of a unimodal relationship between productivity and plant species richness”. Science 351:457CrossRefPubMedGoogle Scholar
  68. Vetaas OR, Grytnes JA (2002) Distribution of vascular plant species richness and endemic richness along the Himalayan elevation gradient in Nepal. Global Ecol Biogeogr 11:291–301CrossRefGoogle Scholar
  69. Walther BA, Moore JL (2005) The concepts of bias, precision and accuracy, and their use in testing the performance of species richness estimators, with a literature review of estimator performance. Ecography 28:815–829CrossRefGoogle Scholar
  70. Webb TJ, Gaston KJ (2003) On the heritability of geographic range sizes. Am Nat 161:553–566CrossRefPubMedGoogle Scholar
  71. Whittaker RH (1952) A study of summer foliage insect communities in the Great Smoky mountains. Ecol Monogr 22:1–44CrossRefGoogle Scholar
  72. Whittaker RH (1960) Vegetation of the Siskiyou mountains, Oregon and California. Ecol Monogr 30:279–338CrossRefGoogle Scholar
  73. Whittaker RH (1967) Gradient analysis of vegetation. Biol Rev 42:207–264CrossRefPubMedGoogle Scholar
  74. Whittaker RJ, Willis KJ, Field R (2001) Scale and species richness: towards a general, hierarchical theory of species diversity. J Biogeogr 28:453–470CrossRefGoogle Scholar
  75. Wohlgemuth T, Nobis MP, Kienast F, Plattner M (2008) Modelling vascular plant diversity at the landscape scale using systematic samples. J Biogeogr 35:1226–1240CrossRefGoogle Scholar
  76. Wu Y, Colwell RK, Rahbek C, Zhang C, Quan Q, Wang C, Lei F (2013) Explaining the species richness of birds along a subtropical elevational gradient in the Hengduan Mountains. J Biogeogr 40:2310–2323CrossRefGoogle Scholar
  77. Zhang Z, He JS, Li J, Tang Z (2015) Distribution and conservation of threatened plants in China. Biol Conserv 192:454–460CrossRefGoogle Scholar

Copyright information

© The Botanical Society of Japan and Springer Japan 2017

Authors and Affiliations

  • Kumar Manish
    • 1
    • 2
  • Maharaj K. Pandit
    • 1
    • 2
    Email author
  • Yasmeen Telwala
    • 1
    • 2
  • Dinesh C. Nautiyal
    • 2
  • Lian Pin Koh
    • 3
  • Sudha Tiwari
    • 1
    • 2
  1. 1.Department of Environmental StudiesUniversity of DelhiDelhiIndia
  2. 2.Centre for Interdisciplinary Studies of Mountain and Hill EnvironmentUniversity of DelhiDelhiIndia
  3. 3.Environment Institute, and School of Biological SciencesUniversity of AdelaideAdelaideAustralia

Personalised recommendations